Zootaxa 2424: 1–41 (2010) Review, Cladistic Analysis

Review, cladistic analysis and biogeography of Nezara Amyot & Serville (Hemiptera: Pentatomidae)

AUGUSTO FERRARI1, CRISTIANO FELDENS SCHWERTNER1, 3 & JOCELIA GRAZIA2 Laboratório de Entomologia Sistemática, Departamento de Zoologia, Programa de Pós-Graduação em Biologia Animal, Universidade Federal do Rio Grande do Sul, Av. Bento Gonçalves, 9500, Bloco IV, prédio 43435, 91501-970 Porto Alegre, RS, Brazil. E-mail: [email protected] acrosternum@yahoo.com.br [email protected] 1CAPES Fellowship 2CNPq Fellowship 3Current address: Departamento de Ciências Biológicas, Universidade Federal de São Paulo – Campus Diadema, Rua Prof. Artur Riedel, 275, 09972-270, Diadema, SP, Brazil

Table of contents

Abstract ...... 2 Introduction ...... 2 Material and methods ...... 3 Results and discussion ...... 5 The genus Nezara ...... 5 Description of characters ...... 6 Analysis ...... 11 Biogeography ...... 13 Taxonomy ...... 15 Nezara Amyot & Serville, 1843 ...... 15 Key to the species of Nezara (male and female): ...... 16 Nezara viridula (Linnaeus, 1758) ...... 16 Nezara antennata Scott, 1874 ...... 18 Nezara yunnana Zheng, 1982 ...... 18 Nezara similis Freeman, 1940 ...... 20 Nezara mendax Breddin, 1908...... 22 Nezara naspira (Dallas, 1851) ...... 24 Nezara capicola (Westwood, 1837) ...... 27 Nezara orbiculata Distant, 1890 ...... 29 Nezara soror Schouteden, 1905 ...... 31 Nezara immaculata Freeman, 1940 ...... 32 Nezara robusta Distant, 1898 ...... 34 Nezara niamensis (Distant, 1890) ...... 35 Acknowledgments ...... 36 References ...... 36

Accepted by M. Malipatil: 29 Jan. 2010; published: 13 Apr. 2010 1 Abstract

In this paper, the genus Nezara Amyot & Serville is reviewed and a hypothesis of relationships among its species is proposed. Also, diagnoses for the genus and the 12 included species are provided. A key to separate the species is also provided. Of the seven other species assigned to Nezara in the literature, four of them are considered incertae sedis, two are transferred to different genera, and one is considered a junior synonym. The cladistic analysis include 15 taxa and 34 characters. The cladogram was rooted with Carpocoris purpureipennis (DeGeer); outgroup comparison also include species of the sister-groups Aethemenes chloris (Dallas), and Pseudoacrosternum cachani Day. The results showed Nezara as a monophyletic taxon. Biogeographical scenarios of the evolution of Nezara were discussed.

Key words: Nezarini, Pentatominae, key to species, synonymies, Afrotropical region, Oriental region

Introduction

Among phytophagous green stink bugs, species of the genus Nezara Amyot & Serville are medium to large in size, and found mainly in the Afrotropical region (Freeman 1940; Linnavuori 1972, 1982). Only three species are distributed elsewhere: N. viridula (Linnaeus, 1758) (cosmopolitan); N. antennata Scott, 1874 (northern Oriental region and Japan), and N. yunnana Zheng, 1982 (northern Oriental region) (Freeman 1940; Zheng 1982; Rider 2006). As far as is known, species of Nezara are polyphagous, some of them related to crop damage (Panizzi et al. 2000). The southern green stink bug, N. viridula, is a serious pest of several important crops worldwide (McPherson & McPherson 2000; Panizzi et al. 2000), and is one of the most studied pentatomid species (i.e. DeWitt & Godfrey 1972; Todd 1989; Panizzi 1997). Amyot & Serville (1843) proposed Nezara in the group ‘Rhaphigastrides’ to include Cimex smaragdulus Fabricius, 1775 (= viridulus Linnaeus) and Pentatoma marginata Palisot de Beauvois, 1817. Dallas (1851) considered Nezara as a subgenus of Rhaphigaster Laporte, 1833, which was followed by Stål (1858, 1860). Stål (1865) recognized Nezara as a valid genus, however its identity and composition were never well understood during the subsequent years (i.e. Mulsant & Rey 1866; Stål 1872, 1876; Lethierry & Severin 1893). Several species were described or included in the genus during that time, but most of them are currently recognized as synonyms or have been transferred to other genera (i.e. Bergroth 1914; Freeman 1940; Orian 1965; Day 1965). To account for the diversity in the genus, Kirkaldy (1909) recognized in his catalogue six subgenera in Nezara, all of which are now considered to be distinct genera (Bergroth 1914; Freeman 1940; Linnavuori 1972; Thomas & Yonke 1981; Rolston 1983; Grazia & Fortes 1995). The taxonomic limits of Nezara as known today were established by Bergroth (1914). He followed Stål (1876) in considering Acrosternum Fieber as a valid genus, using the morphology of the ostiolar peritreme to distinguish it from Nezara. Therefore, following Bergroth (1914), Nezara contained the species placed by Stål (1876) in his division a of the genus. However, only the revision carried out by Freeman (1940) firmly established the limits of the genus; he recognized 11 species of Nezara and 21 varieties; nine of the varieties were described by him. More recently, Schwertner (2005) tested and corroborated the monophyly of Nezara, and proposed a hypothesis of relationship with other genera of the green stink bugs. Most of the species of Nezara have chromatic polymorphism that were described by many authors (e.g. Linnaeus 1758; Fabricius 1803; Horváth 1889; Schouteden 1910; Leston 1953; Freeman 1940; Kiritani & Yukawa 1963), showing great inter- and intraspecific variability. In Nezara viridula, the relative frequency of the chromatic polymorphic forms is dependant on environment conditions, i.e. photoperiod and temperature (Kiritani & Sasaba 1969; Musolin & Numata 2003). Yukawa & Kiritani (1965) hypothesized that the polymorphic forms in a population of N. viridula are determined by the genetic composition followed by genetic drift and natural selection. The genetic basis of polymorphism in N. viridula was demonstrated recently (Ohno & Alam 1992). Although Poe & Wiens (2000) stated that analyses using polymorphic characters are consistently more accurate, we excluded polymorphic characters from our study for two reasons. First, the number of the polymorphic forms for all the species of Nezara was low in our sample. Second, based on the morphological characters used (i.e. genitalia of both sexes), we did not find any evidence to support the polymorphic forms as valid species or subspecies.